ZNF469 and keratoconus: However, the GWAS showed an unexpected effect direction for SNP rs9938149 in BANP-ZNF469, with the CCT-increasing allele leading to increased risk for keratoconus, implying that, in addition to alternative mechanisms which may be specific for keratoconus, part of the genetic predisposition to keratoconus is mediated through the genes underlying CCT [109].